PHOTOSYNTHESIS AND PHOTOSYNTHESIS-COUPLED RESPIRATION IN NATURAL BIOFILMS QUANTIFIED WITH OXYGEN MICROSENSORS1

Photosynthesis and respiration were analyzed in natural biofilms by use of O₂ microsensors. Depth profiles of gross photosynthesis were obtained from the rate of decrease in O₂ concentration during the first few seconds following extinction of light, and net photosynthesis of the photic zone was calculated from O₂ concentration gradients measured at steady state. Respiration within the photic zone was calculated as the difference between gross and net photosynthesis. Two types of biofilms were investigated: one dominated by diatoms, and one dominated by cyanobacteria. High O₂/CO₂ ratios caused increased respiration especially within the diatom biofilm, which could indicate that photorespiration was a dominant O₂-consuming process. The rate of respiration was constant within both biofilms during the first 4.6 s following extinction of light, even when respiration was stimulated by high O₂/CO₂ ratio. The assumption of a constant rate of respiration during the dark period is an essential one for the determination of gross photosynthetic activity by use of O₂ microsensors. We here present the first evidence to substantiate this assumption. The results strongly suggest that gross photosynthesis as measured by use of O₂ microsensors may include carbon equivalents that are subsequently lost through photorespiration. Computer modeling of photosynthesis profiles measured after 1.1, 1.6, and 2.6 s of dark incubation illustrated how the actual photosynthesis profile could have appeared if it had been possible to do the determination at time 0. Diffusion of O₂ during the up to 4.6-s long dark incubations did not affect gross photosynthetic rate when integrated over all depths, but the apparent vertical distribution of the photosynthetic activity was strongly affected.     

IRON STIMULATION OF PHOTOSYNTHESIS AND NITROGEN FIXATION IN ANABAENA 7120 AND TRICHODESMIUM (CYANOPHYCEAE)1

Iron availability may limit carbon and nitrogen fixation in the oceans. The freshwater cyanobacterium, Anabaena, was used as a laboratory model for the biochemical and physiological effects of iron. Increased iron nutrition, in the range of 10^?8 M to 10^?6 M resulted in increases of approximately four fold in carbon and nitrogen fixation rates. Chlorophyll concentration increased, and the relative amount of in vivo fluorescence was reduced with more iron. Natural samples of Trichodesmium, collected off Barbados and incubated with increased iron for two days, showed similar effects. Trichodesmium responded to iron additions indicating that it may be Fe limited in its natural environment. These responses to iron are consistent with the biochemical roles of iron in photosynthesis and nitrogen fixation. The results are discussed in the geochemical context of the sporadic total iron input to tropical oceans and possible implications to spatial and temporal patterns of productivity.     

ANTARCTIC CYANOBACTERIA: LIGHT,NUTRIENTS, AND PHOTOSYNTHESIS IN THE MICROBIAL MAT ENVIRONMENT1

The microenvironmental and photosynthetic characteristics of Antarctic microbial mats were measured in a series of ponds near McMurdo Sound. As elsewhere in Antarctica, these cold-water benthic communities were dominated by oscillatoriacean cyanobacteria. Despite large variations in mat thickness, surface morphology, and color, all of the communities had a similar pigment organization, with a surface carotenoid-rich layer that overlaid a deep chlorophyll maximum (DCM) enriched in phycocyanin as well as chlorophyll a. Spectroradiometric analyses showed that the DCM population inhabited an orange-red shade environment. In several of the mats, the deep-living trichomes migrated up to the surface of the mat within 2 h in response to a 10-fold decrease in surface irradiance. The euphotic layer of the mats was supersaturated in oxygen and contained ammonium and dissolved reactive phosphorus concentrations in excess of 100 mg N·m^?3 or P·m^?3. Integral photosynthesis by core samples was saturated at low irradiances and varied two- to threefold throughout the continuous 24-h radiation cycle. Oxygen microelectrode analyses showed that the photosynthetic rates were slow to negligible near the surface and maximal in the DCM. These compressed, nutrient-rich euphotic zones have some properties analogous to planktonic systems, but the integrated photosynthetic responses of the community reflect the strong self-shading within the mat and physiological dominance by the motile, DCM populations.     

NONEQUILIBRIUM RATES OF PHOTOSYNTHESIS AND RESPIRATION UNDER DYNAMIC LIGHT SUPPLY1

To identify processes that might account for differences in growth rates of rhodophytes under constant and dynamic light supply, we examined nonequilibrium gas exchange by measuring time courses of photoinduction, loss of photoinduction, and respiration rates immediately after the light–dark transition. Using the rhodophyte species Palmaria palmata (Huds.) Lamour and Lomentaria articulata (Huds.) Lyngb., we compared the effects of growth-saturating constant photon flux density (PFD) (95 μmol photons · m^?2· s^?1) to those of a dynamic light supply modeled on canopy movements in the intertidal zone (25 μmol photons · m^?2· s^?1 background PFD plus light flecks of 350 μmol photons · m^?2· s^?1, 0.1 Hz). The time required for P. palmata and L. articulata to be fully photoinduced was not affected by the dynamics of light supply. L. articulata required only 6 min of illumination with either fluctuating or constant light to be completely induced compared to 20 min for P. palmata. The latter species also lost photoinduction more rapidly than did L. articulata in the dark. There was no significant decline in photoinduction state for either species at the background PFD. The time courses of respiration after illumination with constant and fluctuating light were significantly different for P. palmata but not for L. articulata when the total photon dose was equal. In general, gas exchange of P. palmata appeared to be particularly sensitive to the temporal distribution of light supply whereas that of L. articulata was sensitive to the amplitude of variations, being photoinhibited at high PFD. These results are discussed in terms of the different mechanisms of inorganic carbon acquisition in the two species.     

CYANOBACTERIAL ASSEMBLAGES IN PERMANENT ICE COVERS ON ANTARCTIC LAKES: DISTRIBUTION, GROWTH RATE, AND TEMPERATURE RESPONSE OF PHOTOSYNTHESIS

The proliferation of microalgae in the McMurdo Dry Valleys of Antarctica is intricately linked to the seasonal cycle involving the freezing and melting of water. Anecdotal observations and preliminary sampling have found cyanobacterial cells in ice covers on lakes in the McMurdo Dry Valleys, and several of these ice covers are known to undergo seasonal freeze–thaw cycles. Therefore, we sought to determine the distribution and abundance of cyanobacterial assemblages in several permanent ice covers throughout the McMurdo Dry Valleys and to determine their rates of growth and their photosynthetic physiologies upon encountering liquid water. We found that the majority of the permanent ice covers contained cyanobacterial assemblages in close association with sedimentary material. Cyanobacterial biomass was conspicuously absent in sediment-free ice covers, suggesting that the seasonal interaction between the sediments, ice, and solar radiation present the necessary liquid water environment for cyanobacterial growth. All assemblages exhibited extremely low rates of photosynthesis when first exposed to liquid water. Despite the low rates of photosynthesis, a large proportion (41%) of the photosynthate was incorporated into protein, indicating that the cells were undergoing efficient net cellular growth. The short-term response (24 h) of photosynthesis to a range of temperatures showed optimum rates occurring at temperatures >15° C, which is similar to those of psychrotrophic cyanobacteria isolates from soil and stream habitats, which we believe provides the inoculum for the in- ice habitats.     

TOXICITY OF BISULFITE TO PHOTOSYNTHESIS AND RESPIRATION1

Inhabition of photosynthesis in Chloroccoum sp by bisulfileion was the reciprocal of the light intensity curve. Respiration was least affected of the bisulfite after endogenous substrate was reduced by incubation in darkness. Maximum areduction in growth occurred with bisulfile treatment at or above optimal growth temperatures. Maximum phytotoxicity correlated with conditions resulting in maximum metabolic activity. The order of toxicity was –H₂SO₃HSO₃^?SO₃.     

APPARENT LIGHT REQUIREMENT FOR ACTIVATION OF PHOTOSYNTHESIS UPON REHYDRATION OF DESICCATED BEACHROCK MICROBIAL MATS1

Photosynthetic electron transport of beachrock microbial mats growing in the intertidal zone of Heron Island (Great Barrier Reef, Australia) was investigated with a pulse amplitude modulation chl fluorometer providing four different excitation wavelengths for preferential excitation of the major algal groups (cyanobacteria, green algae, diatoms/dinoflagellates). A new type of fiberoptic emitter‐detector unit (PHYTO‐EDF) was used to measure chl fluorescence at the sample surface. Fluorescence signals mainly originated from cyanobacteria, which could be almost selectively assessed by 640‐nm excitation. Even after desiccation for long time periods under full sunlight, beachrock showed rapid recovery of photosynthesis after rehydration in the light (t_1/2～ 15 min). However, when rehydrated in the dark, the quantum yield of energy conversion of PSII remained zero over extended periods of time. Parallel measurements of O₂ concentration with an oxygen microoptode revealed zero oxygen concentration in the surface layer of rehydrated beachrock in the dark. Upon illumination, O₂ concentration increased in parallel with PSII quantum yield and decreased again to zero in the dark. It is proposed that oxygen is required for preventing complete dark reduction of the PSII acceptor pools via the NADPH‐dehydrogenase/chlororespiration pathway. This hypothesis is supported by the observation that PSII quantum yield could be partially induced in the dark by flushing with molecular oxygen. Abbreviations: EDF, emitter‐detector unit; F_o, fluor‐escence yield of dark‐adapted sample; F_m, maximal fluorescence yield measured during saturation pulse; F_v, variable fluorescence yield; LED, light‐emitting diode; PAM, pulse amplitude modulation; PQ, plastoquinone     

EFFECTS OF LIGHT AND TEMPERATURE ON NET PHOTOSYNTHESIS AND DARK RESPIRATION OF GAMETOPHYTES AND EMBRYONIC SPOROPHYTES OF MACROCYSTIS PYRIFERA1

The rates of net photosynthesis as a function of irradiance and temperature were determined for gametophytes and embryonic sporophytes of the kelp, Macrocystis pyrifera (L.) C. Ag. Gametophytes exhibited higher net photosynthetic rates based on oxygen and pH measurements than their derived embryonic sporophytes, but reached light saturation at comparable irradiance levels. The net photosynthesis of gametophytes reached a maximum of 66.4 mg O₂ g dry wt^?1 h^?1 (86.5 mg CO₂ g dry wt^?1 h^?1), a value approximately seven times the rate reported previously for the adult sporophyte blades. Gametophytes were light saturated at 70 μE m^?2 s^?1 and exhibited a significant decline in photosynthetic performance at irradiances 140 μE m^?1 s^?1. Embryonic sporophytes revealed a maximum photosynthetic capacity of 20.6 mg O₂ g dry wt^?1 h^?1 (25.3 mg CO₂ g dry wt^?1 h^?1), a rate about twice that reported for adult sporophyte blades. Embryonic sporophytes also became light saturated at 70 μE m^?2 s^?1, but unlike their parental gametophytes, failed to exhibit lesser photosynthetic rates at the highest irradiance levels studied; light compensation occurred at 2.8 μE m^?2 s^?1. Light-saturated net photosynthetic rates of gametophytes and embryonic sporophytes varied significantly with temperature. Gametophytes exhibited maximal photosynthesis at 15° to 20° C, whereas embryonic sporophytes maintained comparable rates between 10° and 20° C. Both gametophytes and embryonic sporophytes declined in photosynthetic capacity at 30° C. Dark respiration of gametophytes was uniform from 10° to 25° C, but increased six-fold at 30° C; the rates for embryonic sporophytes were comparable over the entire range of temperatures examined. The broader light and temperature tolerances of the embryonic sporophytes suggest that this stage in the life history of M. pyrifera is well suited for the subtidal benthic environment and for the conditions in the upper levels of the water column.     

THE EFFECT OF IRON NUTRITION ON PHOTOSYNTHESIS AND NITROGEN FIXATION IN CULTURES OF TRICHODESMIUM (CYANOPHYCEAE)1

Cultures of Trichodesmium NIBB 1067 were grown in the synthetic medium AQUIL with a range of iron added from none to 5 × 10^?7 M Fe for 15 days. Chlorophyll-a, cell counts, and total cell volume were two or three times higher in medium with 10^?7 M Fe than with no added Fe. Oxygen production rate per chlorophyll-a was over 60% higher with higher iron. Increased iron stimulated photosynthesis at all irradiances from about 12–250 μE · m^?2· s^?1. Nitrogen fixation rate, estimated from acetylene reduction, for 10^?7 and 10^?8 M Fe cultures was approximately twice that of the cultures with no added Fe. The range of rates of O₂ production and N₂ fixation in cultures at the iron concentrations we used were similar to the rates from natural samples of Trichodesmium from both the Atlantic, and the Pacific oceans. This similarity may allow this clone to be used, with some caution, for future physiological ecology studies. This study demonstrates the importance of iron to photosynthesis and nitrogen fixation and suggests that Trichodesmium plays a central role in the biogeochemical cycles of iron, carbon and nitrogen.     

CHANGES IN PHOTOSYNTHESIS IN RESPONSE TO COMBINED IRRADIANCE AND TEMPERATURE STRESS IN CYANOBACTERIAL SURFACE WATERBLOOMS1

Buoyant cyanobacteria, previously mixed throughout the water column, float to the lake surface and form a surface waterbloom when mixing subsides. At the surface, the cells are exposed to full sunlight, and this abrupt change in photon irradiance may induce photoinhibition; at the same time, temperature rises as well. This study investigated the damaging effects of this increase in temperature as well as the ecologically more relevant combination of both an increased temperature and a high photon irradiance. Analysis of surface blooms with oxygen microelectrodes showed that integrated oxygen contents that are dependent on the balance of photosynthetic oxygen evolution and respiratory oxygen uptake decreased when temperature was raised above the lake temperature. Gross rates of photosynthesis were unaffected by temperatures up to of 35°C; hence, a moderate increase in temperature mainly stimulated oxygen uptake. Preincubation of cells of the cyanobacterium Anabaena flos-aquae (Lyngb.) de Brébisson at temperatures up to 35°C did not affect the subsequent measurement of rates of net photosynthesis. Another 5°C rise in temperature severely damaged the photosynthetic apparatus. Failure to restore net rates of photosynthesis was coupled to a strong quenching of the ratio of variable to maximum fluorescence, F_v/F_m, that was the result of a rise in F_o. A combination of high temperature and high photon irradiance was more damaging than high temperature alone. In contrast, low photon irradiances offered substantial protection against heat injury of the photosynthetic apparatus. I conclude from this study that because cyanobacteria usually are acclimated to low average irradiance prior to bloom formation, there is a reasonable risk of chronic photoinhibition. The increase in temperature will enhance the photodamage of cells in the top layer of the bloom. Low photon irradiances in subsurface layers will offer protection against heat injury. If the high temperatures extend to the deepest, dark layers of the bloom, damage in those layers is likely to occur.     

EFFECT OF TEMPERATURE,LIGHT AND pH ON GROWTH,PHOTOSYNTHESIS AND RESPIRATION OF THE DINOFLAGELLATE PERIDINIUM CINCTUM FA. WESTII IN LABORATORY CULTURES1

Optimum light, temperature, and pH conditions for growth, photosynthetic, and respiratory activities of Peridinium cinctum fa. westii (Lemm.) Lef were investigated by using axenic clones in batch cultures. The results are discussed and compared with data from Lake Kinneret (Israel) where it produces heavy blooms in spring. Highest biomass development and growth rates occurred at ca. 23° C and ≥50 μE· m^?2·s¹ of fluorescent light with energy peaks at 440–575 and 665 nm. Photosynthetic oxygen release was more efficient in filtered light of blue (BG 12) and red (RG 2) than in green (VG 9) qualities. Photosynthetic oxygen production occurred at temperatures ranging from 5° to 32° C in white fluorescent light from 10 to 105 μE·m^?2·s^?1 with a gross maximum value of 1500 × 10^?12 g·cell^?1·h^?1 at the highest irradiance. The average respiration amounted to ca. 12% of the gross production and reached a maximum value of ca. 270·10^?12 g·cell^?1·h^?1 at 31° C. A comparison of photosynthetic and respiratory Q₁₀-values showed that in the upper temperature range the increase in gross production was only a third of the corresponding increase in respiration, although the gross production was at maximum. Short intermittent periods of dark (>7 min) before high light exposures from a halogen lamp greatly increased oxygen production. Depending on the physiological status of the alga, light saturation values were reached at 500–1000 μE·m^?2·s^?1 of halogen light with compensation points at 20–40 μE·m^?2·s^?1 and I_k-values at 100–200 μE·m^?2·s^?1. The corresponding values in fluorescent light in which it was cultured and adapted, were 25 to 75% lower indicating the ability of the alga to efficiently utilize varying light conditions, if the adaptation time is sufficient. Carbon fixation was most efficient at ca. pH 7, but the growth rates and biomass development were highest at pH 8.3.     

MICROENVIRONMENTS AND MICROSCALE PRODUCTIVITY OF CYANOBACTERIAL DESERT CRUSTS1

We used microsensors to characterize physicochemical microenvironments and photosynthesis occurring immediately after water saturation in two desert soil crusts from southeastern Utah, which were formed by the cyanobacteria Microcoleus vaginatus Gomont, Nostoc spp., and Scytonema sp. The light fields within the crusts presented steep vertical gradients in magnitude and spectral composition. Near-surface light-trapping zones were formed due to the scattering nature of the sand particles, but strong light attenuation resulted in euphotic zones only ca. 1 mm deep, which were progressively enriched in longer wavelengths with depth. Rates of gross photosynthesis (3.4–9.4 mmol O₂·m^?2·h^?1) and dark respiration (0.81–3.1 mmol O^?2·m^?2·h^?1) occurring within 1 to several mm from the surface were high enough to drive the formation of marked oxygen microenvironments that ranged from oxygen supersaturation to anoxia. The photosynthetic activity also resulted in localized pH values in excess of 10, 2–3 units above the soil pH. Differences in metabolic parameters and community structure between two types of crusts were consistent with a successional pattern, which could be partially explained on the basis of the microenvironments. We discuss the significance of high metabolic rates and the formation of microenvironments for the ecology of desert crusts, as well as the advantages and limitations of microsensor-based methods for crust investigation.     

EFFECTS OF SALINITY AND LIGHT INTENSITY ON THE RESUMPTION OF PHOTOSYNTHESIS IN REHYDRATED CYANOBACTERIAL MATS FROM BAJA CALIFORNIA SUR,MEXICO1

Lyngbya mats in the intertidal of the Laguna Ojo de Liebre are metabolically active for only a few days every 2 weeks during spring tides, with environmental conditions varying greatly during these periods of hydration. Pulse amplitude modulated fluorometry (PAM) and oxygen measurements were used to measure photosynthetic activity during the first few hours after rehydration under various light intensities and salinities. Upon rehydration, a transitory maximum in respiratory activity (10–30 min) occurred before the resumption of photosynthesis, which could recover in about 2 h. Salinities outside the mats' natural range (35–50 psu) were detrimental to photosynthetic recovery. Both high (100 psu) and low (0–10 psu) salinities slowed recovery as well as lowered the overall photosynthetic yield. Photosynthesis was initiated earlier and recovered more rapidly with increasing light intensity. In addition, the positive effect of light on rates of recovery was disproportionately greater at lower salinities (10–25 psu) where high light (500 W·m^?2) counteracted the negative effects of low‐salt stress early in recovery. However, higher light intensities became photoinhibitory later in recovery (>2 h). Photosynthesis did not recover uniformly within the mat. Filaments deeper in the mat most likely recovered later than those near the surface due to high light attenuation. The ability of the mats to tolerate desiccation and take advantage of hydration periods even when conditions are suboptimal enables these mats to predominate in the intertidal environment.     

MICROSENSOR STUDIES OF OXYGEN AND LIGHT DISTRIBUTION IN THE GREEN MACROALGA CODIUM FRAGILE1

Scalar irradiance, oxygen concentration, and oxygenic photosynthesis were measured at 0.1 mm spatial resolution within the tissue of the siphonous green macroalga Codium fragile subsp. tomentosoides (van Goor) Silva by fiber-optic scalar irradiance microsensors and oxygen microelectrodes. The scalar irradiance of visible light was strongly attenuated in the outer 0.2 mm of the tissue but was nearly constant for the subsequent 1.0 mm of photo-synthetic tissue. Far-red scalar irradiance at 750 nm increased below the tissue surface to a maximum of 200% of incident irradiance at 1.2 mm depth due to multiple scattering in the medullary tissue. The constant intensity of visible light below 0.2 mm was thus a result of the combined effects of absorption and backscattering from the medulla. The oxygen exchange between the alga and the surrounding water was diffusion-limited with a steep O₂gradient inside and around the alga. In darkness, the tissue below 0.6 mm became anoxic, and endophytic extracellular space provided an environment where anoxygenic microbial processes may occur. When illuminated at 160 nmol photons·^?2·^?1, O₂ concentrations exceeded ambient levels throughout the thallus, with a maximum of 250% of air saturation just below the surface. The amplitude of oxygen variation was buffered by gas bubbles formed in the medullary tissue.     

CARBON ALLOCATION IN MACROCYSTIS PYRIFERA (PHAEOPHYTA): INTRINSIC VARIABILITY IN PHOTOSYNTHESIS AND RESPIRATION1

Measurements of net photosynthesis (PS, O₂ evolution), dark respiration (R, O₂ consumption), and light and dark carbon fixation (¹⁴C) were conducted on whole blades, isolated blade discs, sporophylls, apical scimitars and representative portions of stipe and holdfast of the giant kelp Macrocystis pyrifera L.C. Ag. On a dry weight basis, highest net PS rates were observed in apical scimitar segments and whole blades (3.81 and 3.07 mgC · g dry wt^?1· h^?1, respectively), followed by sporophylls (1.42 mgC·g dry wt^?1· h^?1) and stipe segments (0.15 mgC·g dry wt^?1· h^?1). No PS capacity was observed in holdfast material. Respiration rates showed similar ranking ranging from 1.22 mgC·g dry wt^?1·h^?1 for apical scimitar to 0.18–0.22 mgC·g dry wt^?1· h^?1 for holdfast material. Considerable within blade variability in both PS and R was also found. Steepest PS and R gradients on both an areal and weight basis were found within immature blades followed by senescent and mature blade material. Highest net PS rates were associated with the blade tips ranging from 3.08 (mature blades) to 10.3 mgC·dry wt^?1·h^?1 (immature blades). Highest rates of R generally occurred towards the basal portions of blades and ranged from 1.03–1.80 mgC·g dry wt^?1·h^?1 for immature blades. The variability within and between blades was high, with coefficients of variation approaching 50%. The observed patterns can be related to the decreasing proportionment of photosynthetic tissue and increasing proportionment of structural tissue as occurs from the blade tip to the blade base. Rates of light carbon fixation (LCF) revealed longitudinal profiles similar to oxygen measurements for the different blade types, with the absolute rates being slightly lower. Patterns of dark carbon fixation (DCF) were less easily interpreted. Highest rates of DCF (0.04–0.06 mgC·g dry wt^?1·h^?1) occurred at the basal portions of immature and senescent blades. Longitudinal profiles of total chlorophyll (a + c) on both an areal and weight basis were very similar to the profiles of PS. Normalized to chlorophyll a, PS displayed an unusual longitudinal profile in immature tissue; however, such profiles for mature and senescent tissues were similar to those for PS on an areal basis. It was demonstrated that it is difficult, if not impossible, to select single tissue discs that are representative of whole blades. The metabolic longitudinal profiles reveal a characteristic developmental pattern; the previous working definitions of immature, mature, and senescent blades, based on morphology and frond position thus have a physiological basis.     

SHORT‐TERM EFFECTS OF ACETATE AND MICROAEROBIC CONDITIONS ON PHOTOSYNTHESIS AND RESPIRATION IN CHLORELLA SOROKINIANA GXNN 01 (CHLOROPHYTA)1

The culture of microalgae using organic carbon sources decreases the cost of operation in closed systems. The effect of carbon sources on microalgae is thus an interesting problem in not only theoretical research but also practical production. The short‐term effects of acetate and microaerobic conditions on the growth, photosynthesis, and respiration of the green microalga Chlorella sorokiniana I. Shihira & R.W. Krauss GXNN 01 were described after acetate addition to autotrophic cultures. As the acetate concentration increased, cells needed a longer lag phase to grow, and 243.8 mM acetate completely inhibited growth. Acetate addition induced an immediate response in photosynthesis and respiration. The activity of PS II and PS I were impaired and declined with different rates, and then recovered compared with autotrophic cells. Carbonic anhydrase and Rubisco activities were also inhibited at the beginning, and respiration was increased. We propose that ATP consumption for acetate assimilation results in surplus NADPH, and then accumulated reducing power over‐reduces inter‐photosystem components and raises the transthylakoid proton gradient, which redistributes energy between PS I and PS II, and leads to a decrease in the PS II/PS I ratio and O₂ evolution. An apparent cyclic electron flow was also observed, which may be mainly mediated by NAD(P)H dehydrogenase‐dependent pathway since NADPH was in excess. These observations pointed to an acclimation process after acetate addition, and suggested the interaction between photosynthesis and respiration involving ATP and reducing power.     

PHOTOACCLIMATION OF PHOTOSYNTHESIS IRRADIANCE RESPONSE CURVES AND PHOTOSYNTHETIC PIGMENTS IN MICROALGAE AND CYANOBACTERIA1

The photosynthesis‐irradiance response (PE) curve, in which mass‐specific photosynthetic rates are plotted versus irradiance, is commonly used to characterize photoacclimation. The interpretation of PE curves depends critically on the currency in which mass is expressed. Normalizing the light‐limited rate to chl a yields the chl a‐specific initial slope (α^chl). This is proportional to the light absorption coefficient (a^chl), the proportionality factor being the photon efficiency of photosynthesis (φ_m). Thus, α^chl is the product of a^chl and φ_m. In microalgae α^chl typically shows little (<20%) phenotypic variability because declines of φ_m under conditions of high‐light stress are accompanied by increases of a^chl. The variation of α^chl among species is dominated by changes in a^chl due to differences in pigment complement and pigment packaging. In contrast to the microalgae, α^chl declines as irradiance increases in the cyanobacteria where phycobiliproteins dominate light absorption because of plasticity in the phycobiliprotein:chl a ratio. By definition, light‐saturated photosynthesis (P_m) is limited by a factor other than the rate of light absorption. Normalizing P_m to organic carbon concentration to obtain P_m^C allows a direct comparison with growth rates. Within species, P_m^C is independent of growth irradiance. Among species, P_m^C covaries with the resource‐saturated growth rate. The chl a:C ratio is a key physiological variable because the appropriate currencies for normalizing light‐limited and light‐saturated photosynthetic rates are, respectively, chl a and carbon. Typically, chl a:C is reduced to about 40% of its maximum value at an irradiance that supports 50% of the species‐specific maximum growth rate and light‐harvesting accessory pigments show similar or greater declines. In the steady state, this down‐regulation of pigment content prevents microalgae and cyanobacteria from maximizing photosynthetic rates throughout the light‐limited region for growth. The reason for down‐regulation of light harvesting, and therefore loss of potential photosynthetic gain at moderately limiting irradiances, is unknown. However, it is clear that maximizing the rate of photosynthetic carbon assimilation is not the only criterion governing photoacclimation.     

KINETICS OF BLUE-LIGHT STIMULATION AND CIRCADIAN RHYTHMICITY OF LIGHT-SATURATED PHOTOSYNTHESIS IN BROWN ALGAE: A SPECIES COMPARISON1

The time courses of photosynthetic rates in red light, with and without additional blue light, were investigated and compared in 20 species of brown algae. Species could be separated into two groups on the basis of the rhythmicity of their photosynthesis in red light and the kinetics of their responses to blue-light pulses. One group, which consisted of members of the Ectocarpales, Chordariales, and Dictyosiphonales, was characterized by strong and persistent circadian rhythmicity in red light. The photosynthetic responses of these species to blue-light pulses started within 10–30 s of the beginning of blue-light treatment and mostly contained at least two distinct kinetic components. An early component, which reached a maximum about 5–10 min after the blue-light pulse, was always detectable. Later components were seen as separate peaks or shoulders after an additional 10–20 min. The decay of the response in this group of species was mostly slow, with half-lives of between 0.5 and 1.5 h. In the second group of species, consisting of members of the Dictyotales, Laminariales, and Fucales, photosynthesis in red light was usually non-rhythmic, although circadian rhythms with a weak amplitude or of transient occurrence were observed in some plants of some species. The increase in photosynthesis in response to a blue-light pulse was not detectable until 70–330 s after the start of blue-light treatment, and the response itself had only a single component, with a maximum after about 10 min and half-life of 10–20 min. The lengths of the lag-phases were positively correlated with the times taken to reach the peak in this group, although the lag-phases and the half lives sometimes varied with time in individual plants. Two members of the Sphacelariales (Sphacelaria, Cladostephus) did not fit into either of the two groups because their photosynthesis was rhythmic, but their responses had long lag-phases, a single component, and moderately long half-lives. The differences in the kinetics of the photosynthetic response to blue-light pulses, which have been described for the two main groups of species, are thought to indicate that there are two distinct mechanisms by which light-saturated photosynthesis responds to blue light in brown algae. Since in some species the maximal photosynthesis after a blue-light pulse and the rate of photosynthesis in continuous blue light also varied in a circadian pattern, the response to blue light itself may be under circadian control.     

ABSORPTION AND UTILIZATION OF IRRADIANCE BY CYANOBACTERIAL MATS IN TWO ICE-COVERED ANTARCTIC LAKES WITH CONTRASTING LIGHT CLIMATES

We investigated the under-ice light climate and the efficiency with which light was absorbed and utilized by benthic algal mats in Lakes Hoare and Vanda, two perennially ice-covered lakes in the McMurdo Dry Valleys area of Southern Victoria Land, Antarctica. The ice cover and water column of Lake Vanda were much more transparent than those of Lake Hoare (18% vs. 2% transmission though ice and attenuation coefficients for downwelling irradiance of 0.05 vs. 0.12 m ^{− 1}, respectively). In both lakes the under-ice spectra were dominated by blue-green wavelengths. The benthic flora under perennial ice covers of both lakes comprised thick mucilaginous mats, dominated by cyanobacteria. The mats were well suited to absorb the dominant blue-green wavelengths of the under-ice light, with phycoerythrin being present at high concentrations. The pigment systems of the benthic mats absorbed 30%–50% of the light that reached them, varying with depth and lake. There was a tendency for the percentage of absorption to increase as ambient irradiance decreased. The efficiency of utilization of absorbed irradiance was examined by constructing absorbed irradiance/oxygen evolution curves to estimate community quantum yield. Mats from 13 m in Lake Hoare showed the highest quantum yields, approaching 1 mol of carbon fixed for every 8 mol quanta absorbed under light-limiting conditions. Lake Vanda mats had lower quantum yields, but these increased with depth. Calculated in situ irradiance occasionally exceeded the measured saturating irradiance for oxygen evolution in both lakes, thus efficiency in situ was below the maximum at times. As in other environments, optimization strategies allowed efficient capture and utilization of the lower and middle ranges of experienced irradiance but led to a compromised capacity to use the highest irradiances encountered at each depth.     

EFFECTS OF PHOTON FLUENCE RATE AND LIGHT SPECTRUM COMPOSITION ON GROWTH,PHOTOSYNTHESIS AND PIGMENT RELATIONS IN GRACILARIA SP.1

The optimal photon fluence rate for growth of tha llus tips of Gracilaria sp. was low (about 100 μE·^–2·¹); higher photon fluence rates inhibited growth. Both phycoerythrin (PE) and chlorophyll (chl) contents decreased with increasing photon fluence rates (up to 100 μE·–m^–2s^–1) in a fashion inverse to the growth response. Chl/PE ratios varied directly as the growth response over a larger photon fluence rate range. The peak chl/PE ratios were obtained at a photon fluence rate optimal for growth, suggesting that this parameter may be used to estimate in situ growth rates. A low compensation point (about 7 μE·^–2s^–1) was observed for low light (15 μE·^–2s^–1) grown plants. This compensation point was also obtained for growth in the long–term (5–6 weeks) experiments. Plants grown at 60 and 140 μE·^–2s^–1 showed higher light compensation and saturation points, suggesting that the variations in pigment composition found between the different treatments determine the photosynthetic responses at sub–optimal photon fluence rates. Photosynthetic rates at light saturation were the same, on a biomass basis, for plants grown at the various photon fluence rates. Thus, the photosynthetic dark reactions were not influenced by previous light regimes. It is suggested that maximal photosynthetic rates expressed on a biomass basis better reflect the potential productivity at tight saturation than if expressed on a pigment basis. Gracilaria sp. grew better under non–filtered fluorescent and greenish than under reddish and blue–enriched light of equal and sub–optimal photon, fluence rate. However, the pigment relations of the algae did not change in a direction complementary to the light composition at which they grew. This, together with the relatively higher photosynthetic rates under reddish and blueish light for plants previously grown under reddish and blueish light, suggests that adaptations to variouslight spectra are based on mechanisms different from complementary chromatic adaptation of the pigments.